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The Morphology of the Male and Female Reproductive ZOBODAT - www.zobodat.at Zoologisch-Botanische Datenbank/Zoological-Botanical Database Digitale Literatur/Digital Literature Zeitschrift/Journal: Arthropod Systematics and Phylogeny Jahr/Year: 2017 Band/Volume: 75 Autor(en)/Author(s): Kienbaum Katja, Scholtz Gerhard, Becker Carola Artikel/Article: The morphology of the male and female reproductive system in two species of spider crabs (Decapoda: Brachyura: Majoidea) and the issue of the velum in majoid reproduction 245-260 75 (2): 245– 260 8.9.2017 © Senckenberg Gesellschaft für Naturforschung, 2017. The morphology of the male and female reproductive system in two species of spider crabs (Decapoda: Brachyura: Majoidea) and the issue of the velum in majoid reproduction Katja Kienbaum *, §, 1, Gerhard Scholtz 1, 2 & Carola Becker 1, 2, 3 1 Humboldt-Universität zu Berlin, Institut für Biologie, Vergleichende Zoologie, Philippstr. 13, 10115 Berlin, Germany; Katja Kienbaum * [katja. [email protected]]; Gerhard Scholtz [[email protected]]; Carola Becker [[email protected]] — 2 Cluster of Excellence “Image Knowledge Gestaltung”, Humboldt-Universität zu Berlin, Sophienstr. 22a, 10178 Berlin, Germany — 3 Queen’s University Marine Laboratory; 12–13 The Strand, Portaferry, Co. Down; BT22 1PF, UK — § Former name ‘Katja Jaszkowiak’; * Corresponding author Accepted 04.iv.2017. Published online at www.senckenberg.de/arthropod-systematics on 30.viii.2017. Editors in charge: Stefan Richter & Klaus-Dieter Klass Abstract The reproductive system of spider crabs (Majoidea) has raised considerable interest due to the complexity of female sperm storage or- gans. In several majoid species, the seminal receptacle has been described as being divided into a dorsal storage chamber and a ventral fertilization chamber separated by a muscular velum. The velum is supposed to control the amount of sperm used for fertilization and to play an important role in sperm competition. Here, we present a study on the reproductive systems of the two majoid species, Mithraculus sculptus (Lamarck, 1818) and Stenorhynchus seticornis (Herbst, 1788) using various morphological techniques such as µCT scans and 3D-reconstructions, complemented by paraffin histology. The male gonopods of the herein investigated species are similar in their general morpho logy and in the presence and distribution of setae. The tubular first gonopod holding the ejaculatory canal is much longer than the short and stout second gonopod, which is supposed to function as a piston in the transport of sperm into the female ducts. The female reproductive system of M. sculptus and S. seticornis conforms to that of other Eubrachyura in possessing paired ovaries, oviducts, seminal receptacles, vaginae, and vulvae. Based on our 3D-reconstructions we demonstrate that there is no division of the seminal receptacle into two chambers separated by a velum. In contrast to this, we observed a spatially restricted invagination of the seminal receptacle. A com- parison of our data with those of previous studies, allows for the conclusion that the invagination of the seminal receptacle may have been misinterpreted and mistaken for a velum by other authors. Thus, the division of the seminal receptacle into two chambers separated by a velum is a character which needs to be re-evaluated. Key words Seminal receptacle, storage chamber, insemination chamber, gonopods, setae, 3D-reconstruction. 1. Introduction The Majoidea or spider crabs is a very diverse brachy- al. 1992; PORTER et al. 2005). However, data concerning uran group, comprising more than 800 species, with a the phylogenetic position of the Majoidea within the Eu- worldwide distribution in marine waters (DE GRAVE et brachyura is sparse and partially contradictory (BRÖSING al. 2009). It has been proposed to be a basal branching et al. 2006; TSANG et al. 2014). Whilst their monophyly is lineage within the Eubrachyura based on morphologi- widely accepted (HULTGREN & STACHOWICZ 2008; MAHON cal (JAMIESON et al. 1995) and molecular data (SPEARS et & NEIGEL 2008; TSANG et al. 2014), relationships among ISSN 1863-7221 (print) | eISSN 1864-8312 (online) 245 Kienbaum et al.: The reproductive system of Majoidea the majoid groups remain disputed. The only constant is construction, together with established histological tools, the monophyly of the Oregoniidae and their position as our results reveal that seminal receptacles of both studied sister group to the remaining Majoidea (see HULTGREN et species are not divided into two separate chambers and al. 2009 and GUINOT et al. 2013 for review). The mor- shed doubt on previous interpretations of such a division phology of brachyuran reproductive systems has been and the presence of a velum (DIESEL 1989; SAL MOYANO et discussed with respect to phylogenetic (GUINOT 1977; al. 2010; GONZÁLEZ-PISANI et al. 2011). The present study MCLAY & LÓPEZ GRECO 2011; GUINOT et al. 2013; BECKER demonstrates the value of 3D-reconstructions to under- & SCHOLTZ 2017) and functional analyses (DIESEL 1989; stand the spatial organization of reproductive systems BENINGER et al. 1991). and the need for further studies in order to re-evaluate the The male copulatory system is a complex arrange- majoid reproductive systems. ment of three paired parts, consisting of the first and sec- ond gonopods and the penes. The tubular first gonopod (G1) forms the ejaculatory canal with a basal and distal opening. During copulation, the second gonopod (G2) 2. Materials & methods and the penis are inserted into the G1 through the basal opening. The penis extrudes the sperm into the ejacula- tory canal of the G1. It is then pushed by the G2 into 2.1. Material the female genital duct (BAUER 1986). Gonopods contain valuable phylogenetic information (BENINGER et al. 1991; The specimens of Stenorhynchus seticornis and Mithra- BENINGER & LAROCQUE 1998). This is based on their di- culus sculptus were obtained from commercial vendors verse and specific morphology including the patterns of (www.shop-meeresaquaristik.de). Three females of each setae in combination with the conservative nature of go- species were used for histology and the 3D-reconstruc- nopods (MARTIN & ABELE 1986; VALLINA et al. 2014). tion of the reproductive organs. All investigated females The female reproductive system of eubrachyuran crabs were mature and two females of M. sculptus were oviger- consists of paired ovaries, oviducts, seminal receptacles, ous. vaginae, and vulvae. The seminal receptacles play a major role in eubrachyuran reproduction due to their ability to store sperm from one or more males that can be used to 2.2. Histology fertilize several consecutive broods (CHEUNG 1968). The reproductive morphology of the Majoidea has For the histological analyses, specimens were cold-an- been addressed in several studies (DIESEL 1991; BENI- aes thetised in a freezer at – 18°C for 15 minutes. Whole NGER et al. 1993; LANTEIGNE et al. 1996; SAINTE-MARIE specimens were preserved either in Bouin’s solution & SAINTE-MARIE 1998; ROTLLANT et al. 2007; GONZÁLEZ- or in “Susa Heidenhain” (MORPHISTO® Evolutions- PISANI et al. 2011; ANTUNES et al. 2016). In contrast to forschung und Anwendung GmbH, Frankfurt am Main, other eubrachyurans, a division of the seminal receptacle Germany) for 48 – 73 hours. For decalcification, speci- into two distinct chambers has been described in some mens were treated in Ethylenediaminetetraacetic acid majoids (DIESEL 1989). According to this view, a mus- (EDTA) for 48 – 72 hours. Specimens were then dehy- cular diaphragm, the velum, separates the seminal re- drated through a series of ethanol solutions and infiltrat- ceptacle into a dorsal secretory “storage chamber” filled ed (Shandon Hypercenter XP, Thermo Fisher Scientific, with sperm and a ventral cuticle “insemination chamber” Waltham, Massachusetts, USA) and embedded with par- with the oviduct junction (DIESEL 1989). The putative affin. Sections were prepared at 6 – 8 µm using a rotary existence of a velum has stimulated inferences on the microtome Leica RM2255 (Leica Microsystems GmbH, reproduction of spider crabs in terms of the location of Wetzlar, Germany). All histological sections were stained fertilization (DIESEL 1991) and sperm competition of eu- with the trichromatic Masson-Goldner “light green” brachyurans in general (MCLAY & LÓPEZ GRECO 2011). (MORPHISTO®, Frankfurt am Main, Germany). To date, it is not clear which majoid groups actually pos- sess a velum and whether all structures referred to as a velum (SAL MOYANO et al. 2010; GONZÁLEZ-PISANI et al. 2.3. Scanning electron microscopy (SEM) 2011) are really similar to its original description and definition (DIESEL 1989). The dissected gonopods of two male specimens of each In this study, we investigate the male and female re- species were cleaned manually after an ultrasonic bath. productive morphology of the majoids Stenorhynchus The first and second gonopods were critical point dried seticornis (Herbst, 1788), belonging to the Inachoididae, (Bal-Tec CPD 030, Balzers, Liechtenstein) and sputter and Mithraculus sculptus (Lamarck, 1818) of the Ma- coated with a gold layer (Bal-Tec SCD 005, Balzers, jidae. Our main goal is to re-evaluate characters of the Liechtenstein). The micrographs were taken using a LEO female reproductive system described by DIESEL (1989, (Zeiss) 1430 scanning electron microscope (Carl Zeiss 1990, 1991), in particular the division of the seminal Nano Technology Systems GmbH, Oberkochen, Germa- receptacle into two chambers. By means of the latest ny) and images were
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